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Ornamentation, Behavior, and Maternal Effects in the Female Northern Cardinal

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Ornamentation, Behavior, and Maternal Effects in the Female Northern Cardinal The University of Southern Mississippi The Aquila Digital Community Master's Theses Summer 8-2011 Ornamentation, Behavior, and Maternal Effects in the Female Northern Cardinal Caitlin Winters University of Southern Mississippi Follow this and additional works at: https://aquila.usm.edu/masters_theses Part of the Biology Commons, and the Ornithology Commons Recommended Citation Winters, Caitlin, "Ornamentation, Behavior, and Maternal Effects in the Female Northern Cardinal" (2011). Master's Theses. 240. https://aquila.usm.edu/masters_theses/240 This Masters Thesis is brought to you for free and open access by The Aquila Digital Community. It has been accepted for inclusion in Master's Theses by an authorized administrator of The Aquila Digital Community. For more information, please contact [email protected]. The University of Southern Mississippi ORNAMENTATION, BEHAVIOR, AND MATERNAL EFFECTS IN THE FEMALE NORTHERN CARDINAL by Caitlin Winters A Thesis Submitted to the Graduate School of The University of Southern Mississippi in Partial Fulfillment of the Requirements for the Degree of Master of Science Approved: _Jodie M. Jawor_____________________ Director _Frank R. Moore_____________________ _Robert H. Diehl_____________________ _Susan A. Siltanen____________________ Dean of the Graduate School August 2011 ABSTRACT ORNAMENTATION, BEHAVIOR, AND MATERNAL EFFECTS IN THE FEMALE NORTHERN CARDINAL by Caitlin Winters August 2011 This study seeks to understand the relationship between ornamentation, maternal effects, and behavior in the female Northern Cardinal (Cardinalis cardinalis). Female birds possess ornaments that indicate a number of important known aspects of quality and are usually costly to maintain. However, the extent to which female specific traits, such as maternal effects, are indicated is less clear. It is predicted by the Good Parent Hypothesis that this information should be displayed through intraspecific signal communication. Specifically, androgens and carotenoids are of interest in this study because both are linked to ornamentation, and are also important egg components that impact offspring quality. Additionally, androgens have implications for adult behavior; testosterone specifically is well known to affect aggression. However, results from this study do not support these hypotheses. Instead, no association was found between ornamentation and maternal effects or behavior. We suggest that it is not profitable for signals to be maintained if the cost of maintenance of such traits is more than the value of the information they display. This research indicates that sexually selected traits should be looked at in the context of the organism’s life history in order to determine functionality. ii ACKNOWLEDGMENTS The writer would like to thank the thesis advisor, Dr. Jodie Jawor, and the other committee members, Dr. Frank Moore and Dr. Robb Diehl, for their advice and support throughout the duration of this project, especially Dr. Jodie Jawor, for invaluable advice and guidance through all phases of this project. Additionally, I would like to thank Dr. Kevin Keuhn for invaluable assistance with his HPLC apparatus and for the generous use of his lab. I would also like to thank Dr. Carl Qualls and Dr. Joel McGlothlin for statistical advice and guidance. A special thanks goes to M. Susan DeVries and Aaron Holbrook for their invaluable assistance in collecting this data and for their generous advice on many matters concerning this project. Additional thanks go to undergraduate assistants Jeff Hooker, Matt Duckworth, Anne Winters, and Megan Winters for assistance with fieldwork. iii TABLE OF CONTENTS ABSTRACT ………………………………………………………………..……………ii ACKNOWLEDGMENTS ……………………..….....…………...…………..…...….....iii LIST OF TABLES ..…………………………………........…………………..………….v LIST OF ILLUSTRATIONS ……………………………………………..……………..vi CHAPTER I. INTRODUCTION: ORNAMENTATION AND THE NORTHERN CARINAL ………………………………………….................………... 1 Hormones Ornament Hypotheses The Northern Cardinal II. ORNAMENTATION AND MATERNAL EFFECTS IN THE FEMALE NORTHERN CARDINAL .…….......................................... 15 Introduction Materials and Methods Results Conclusion III. BEHAVIOR IN THE FEMALE NORTHERN CARDINAL .…........ 37 Introduction Materials and Methods Results Conclusion IV. DISCUSSION .……………………..…….…………………………... 52 APPENDIX .................................................................................................................... 60 WORKS CITED .………………...…………………………………………………… 61 iv LIST OF TABLES Table 1. Showing Descriptive Statistics for Ornaments and Egg Components...................27 2. ANCOVA Model Comparing Dependent Variable Egg Carotenoid Levels to Fixed Factors Egg Number, Bird ID, and Year.....................................................27 3. ANCOVA Model Comparing Dependent Variable Egg Testosterone Levels to Fixed Factors Egg Number, Bird ID, and Year.....................................................28 4. Showing Spearman Rank Correlation Results Comparing Egg Carotenoid Levels, Egg T Levels, Hue/Brightness and Saturation and Mask Brightness.......30 5. Showing Descriptive Statistics for Ornamentation and Behavior.........................44 6. Showing Spearman Rank Correlations between Mask Melanin, Mask Score, Crest Length, and Behavioral Score......................................................................45 7. Showing Spearman Rank Correlations between Hue/B, Saturation, Crest Length, and Behavioral Score................................................................................46 v LIST OF ILLUSTRATIONS Figure 1. Facemasks in Female Northern Cardinals, Showing Variation in Expression and the Scoring System …………………..............................……………........ 13 2. Female Northern Cardinal Carotenoid-Based Underwing Ornament……….…. 14 3. Egg Carotenoid Levels ...................................................................................... 28 4. Egg Testosterone Levels ................................................................................... 29 5. Hue/Brightness vs. Egg Carotenoid .................................................................. 30 6. Saturation vs. Egg Carotenoid .......................................................................... 31 7. Melanin Ornament vs. Egg Testosterone ........................................................... 31 8. Hue/B vs. Egg Testosterone ............................................................................... 32 9. Saturation vs. Egg Testosterone ......................................................................... 32 10. Melanin Ornament vs. Egg Carotenoid ............................................................. 33 11. Egg Testosterone vs. Egg Carotenoid ................................................................ 33 12. Mask Melanin vs. Behavioral Score .................................................................. 45 13. Hue/Brightness vs. Behavioral Score ................................................................ 46 14. Saturation vs. Behavioral Score ......................................................................... 47 15. Mask Melanin vs. Hue/B ................................................................................... 48 vi 1 CHAPTER I INTRODUCTION: ORNAMENTATION AND THE NORTHERN CARDINAL Ornamental plumage, conspicuous colors, and other visual signals in animals have historically been the subject of much study and debate (Andersson 1994). Animals across all taxa commonly display traits that cannot be explained by natural selection. The existence of characteristics that seemingly convey no survival advantage was a quandary for early evolutionary biologists. Wallace (1872), Poulton (1890) and Darwin (1871) all had early competing theories on the relevance and function of these traits. Wallace claimed that conspicuous traits function primarily as warning signals, and did not accept Darwin’s theories that mate choice and competition could act as selective pressure. Poulton erroneously supported Wallace’s opinions on warning coloration and also claimed that bright coloration could evolve as a signal to benefit predators. Both authors addressed part of the problem of conspicuousness in traits by acknowledging that some brightly colored individuals display warning information, but they did not address the problem in full. Primarily, why there is a difference in many conspicuous traits in male and female individuals, why many traits are used in competitions for mates, why only sexually mature organisms usually display full expression of the trait, and why non- toxic organisms display conspicuous or colorful traits. Although much debate remains surrounding mechanisms and functions, Darwin’s theory prevailed and simply states that sexually selected traits are a result of differences in reproductive success caused by competition over mates. Darwin revealed a pattern in certain conspicuous traits that he considered the result of what he termed sexual selection (see Andersson 1994): 1. Traits should not be acquired before sexual maturity. 2 2. Females should develop the trait to a lesser degree than males. 3. Males should only develop or display the trait when mating can occur. 4. The trait is displayed to potential mates or sexual rivals (i.e., members of the same species). Indeed, many of the conspicuous traits seen in animals fit this pattern. The colorful plumage of birds grown during the breeding season, and the antlers grown by male deer during the breeding season are both good examples
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